CONCEPTACLE STRUCTURE OF THE PARASITIC CORALLINE RED ALGA CHOREONEMA THURETII (CORALLINALES) AND ITS TAXONOMIC IMPLICATIONS1

The major diagnostic features for erecting the red algal subfamily Choreonematoideae (Corallinales) were a combination of 1) absence of both cell fusions and secondary pit connections, 2) conceptacle roof and wall comprised of a single cell layer, and 3) presence of tetrasporangial pore plugs within a uniporate conceptacle in the monotypic taxon Choreonema thuretii (Bornet) Schmitz. Because this alga is a parasite, the absence of secondary cell connections is most likely an adaptation to a reduced thallus. This study shows that all conceptacles are not composed of a file of cells but rather a single layer of epithallial cells that are underlain by a thick layer of calcified acellular material; both epithallial cells and the calcified layer are produced by peripheral sterile cells. Although the outermost tetrasporangial pore canal is uniporate, there is a calcified acellular multiporate plate recessed just below the rim. The plate is produced by interspersed sterile cells and is continuous with the calcified layer supporting the conceptacle. These unique structures are likely due to parasitism rather than to the ancestral state. Based on these results and a reexamination of published micrographs depicting lenticular cells in Austrolithon intumescens Harvey et Woelkerling, we propose that both subfamily Choreonematoideae and Austrolithoideae are closely allied with subfamily Melobesioideae. This distant relationship to its host (Corallinoideae) plus a combination of unique conceptacle and unusual type of parasitism indicates that C. thuretii is an alloparasite and that it is likely the most ancient red algal parasite studied to date.     

Mastophoropsis canaliculata (Harvey in Hooker) gen. et comb. nov. (Corallinaceae,Rhodophyta) in Southern Australia

Mastophoropsis canaliculata (Harvey in Hooker) gen. et comb. nov. (Corallinaceae, Rhodophyta) is restricted to south-eastern Australian waters. It is unique among Corallinaceae in possessing an erect, tenacular, branched, taeniform, non-geniculate thallus which produces multiporate tetrasporangial conceptacles. Based on a detailed morphological and anatomical study, including an examination of the designated lectotype, this taxon is referred to the tribe Phymatolitheae in the subfamily Melobesioideae and its relationships to other non-geniculate Corallinaceae are discussed. A simplified microtechnique procedure involving decalcification with nitric acid, resin embedding and staining serially mounted sections with KMnO₄ also is outlined. X-ray microanalysis of surface tissues indicates that calcification occurs largely as CaCO₃ and that various structures contain substantially differing amounts of Ca.     

MOLECULAR SYSTEMATICS OF THE XANTHOPHYCEAE

The Corallinales includes ca. 40 genera of calcified red seaweeds. Species are of two distinct morphotypes; those that possess genicula (uncalcified nodes) and those that lack genicula. Most nongeniculate species take the form of crusts. The presence (or absence) of genicula, secondary pit connections, and tetrasporangial conceptacle features have traditionally been used as key characters for delimiting coralline subfamilies. In this study, nuclear encoded 18S and 26S r RNA gene sequences were determined and used to reexamine relationships among coralline taxa. Separate and combined phylogenetic analyses of these data yielded similar trees in which four major lineages are resolved. Heydrichia and Sporolithon (Sporolithaceae) are positioned at the base of the tree and appear to be distantly related to other species examined. Within the Corallinaceae, the nongeniculate Melobesioideae is resolved as a monophyletic group. All members of this subfamily produce mutiporate tetraspoangial conceptacles. The Corallinoideae, which are characterized by unizonate genicula, are resolved as sister to a clade containing species placed in the Lithophylloideae, Mastophoroideae and Metagoniolithoideae. The molecular data indicate that geniculate and nongeniculate species characterized by the presence of secondary pit connections are closely related. For example, both data sets robustly support a sister taxon relationship between Amphiroa and Titanoderma. Our results indicate that: 1) all taxa in which secondary pit connections are present should be referred to the Lithophylloideae and, 2) genicula are nonhomologous structures that are independently derived in Amphiroa, Lithothrix, Metagoniolithon and the last common ancestor of the Corallinoideae.     

PHYLOGENY AND CLASSIFICATION OF REEF‐BUILDING CORALLINE ALGAE (CORALLINALES,RHODOPHYTA)

The Corallinales includes ca. 40 genera of calcified red seaweeds. Species are of two distinct morphotypes; those that possess genicula (uncalcified nodes) and those that lack genicula. Most nongeniculate species take the form of crusts. The presence (or absence) of genicula, secondary pit connections, and tetrasporangial conceptacle features have traditionally been used as key characters for delimiting coralline subfamilies. In this study, nuclear encoded 18S and 26S rRNA gene sequences were determined and used to reexamine relationships among coralline taxa. Separate and combined phylogenetic analyses of these data yielded similar trees in which four major lineages are resolved. Heydrichia and Sporolithon (Sporolithaceae) are positioned at the base of the tree and appear to be distantly related to other species examined. Within the Corallinaceae, the nongeniculate Melobesioideae is resolved as a monophyletic group. All members of this subfamily produce mutiporate tetraspoangial conceptacles. The Corallinoideae, which are characterized by unizonate genicula, are resolved as sister to a clade containing species placed in the Lithophylloideae, Mastophoroideae and Metagoniolithoideae. The molecular data indicate that geniculate and nongeniculate species characterized by the presence of secondary pit connections are closely related. For example, both data sets robustly support a sister taxon relationship between Amphiroa and Titanoderma. Our results indicate that: 1) all taxa in which secondary pit connections are present should be referred to the Lithophylloideae and, 2) genicula are nonhomologous structures that are independently derived in Amphiroa, Lithothrix, Metagoniolithon and the last common ancestor of the Corallinoideae.     

PARASITIC INTERACTIONS AND VEGETATIVE ULTRASTRUCTURE OF CHOREOAEMA THURETII (CORALLINALES,RHODOPHYTA)1

The monotypic coralline red alga, Choreonema thuretii (Bornet) Schmitz (Choreonematoideae), grows endophytically within three geniculate genera of the Corallinoideae. Although the thallus of Choreonema is reduced, lacks differentiated plastids, and is endophytic except for its conceptacles, its status as a parasite has been questioned because cellular connections to the host had not been ob served. Transmission electron microscopy, however, disclosed a previously undescribed type of parasitic interaction in which Choreonema interacts with its host through specialized cells known as lenticular cells. These small, lens-shaped cells are produced from the single file of host-penetrating vegetative cells. Pit plug morphology between vegetative and lenticular cells is polarized. Plug caps facing the vegetative cell have normal coralline morphology, while those facing the lenticular cell are composed of three layers. Regions of lenticular cells near host cells protrude toward the host cell; upon encountering the host cell wall, the prolrusion produces numerous finger-like fimbriate processes that make cellular connections with the host cell. Lenticular cells may extend several protrusions toward a host cell or penetrate more than one host cell; two or more lenticular cells may also penetrate the same host cell. The lack of secondary pit connections, cell fusions, and passage of parasitic nuclei suggest that this parasitic relationship may be evolutionarily older than previously reported cases of parasitism in red algae.     

Late Hauterivian coralline algae (Rhodophyta,Corallinales) from the Iberian Chain (E Spain). Taxonomy and the evolution of multisporangial reproductive structures

Upper Hauterivian reefal carbonates of the Llàcova Formation (Maestrat Basin, Iberian Chain, E Spain) contain Sporolithon phylloideum (Bucur and Dragastan) Tomás, Aguirre, Braga and Martín-Closas comb. nov. and Sporolithon rude (Lemoine) Ghosh and Maithy (1996). Moussavian et al. (1993) identified them as Parakymalithon phylloideum (Bucur and Dragastan) Moussavian 1987 and Archaeolithothamnium rude Lemoine 1925. The re-assessment of the type of P. phylloideum and additional material indicate that the diagnostic characters of the genus do not warrant separation from Sporolithon and the new combination Sporolithon phylloideum is proposed. The lectotype of Sporolithon rude presents sporangial cavities grouped in sori that can be merged originating a structure that resembles the multiporate tetrasporangial conceptacles of the Hapalidiaceae. We hypothesize that multiporate tetrasporangial conceptacles could have originated from the fusion of several sporangial cavities, suggesting a phylogenetic linkage between Sporolithaceae and Hapalidiaceae supported by other anatomical features, molecular phylogeny and the fossil record.     

A PHYLOGENETIC STUDY OF THE CORALLINALES (RHODOPHYTA) BASED ON NUCLEAR SMALL-SUBUNIT rRNA GENE SEQUENCES

Conflicting classifications for the Corallinales were tested by analyzing partial sequences for the nuclear small-subunit ribosomal RNA (SSU) gene of 35 species of coralline algae. Parsimony and likelihood analyses of these data yielded congruent hypotheses that are inconsistent with classifications for the group that include as many as eight subfamilies. Four major clades are resolved within the order, including the early-diverging Sporolithaceae as well as the Melobesioideae and Corallinoideae. The fourth clade, which is supported robustly, includes both nongeniculate and geniculate species classified in the subfamilies Mastophoroideae, Metagoniolithoideae, Lithophylloideae, and Amphiroideae. Molecular and morphological data support the proposal that the latter two subfamilies are sister taxa. Although relationships among some genera are not resolved clearly, the order of branching of taxa among and within the four principal lineages is concordant with paleontological evidence for the group. Relationships inferred among genera within each of the clades is discussed. Seven morphological characters delimiting higher taxonomic groups within the order were combined with the sequence data, analyzed, and optimized onto the resulting tree(s). Except for the presence or absence of genicula, all other characters were found to be phylogenetically informative. Genicula are nonhomologous structures that evolved independently in the Amphiroideae, Corallinoideae, and Metagoniolithoideae. The phenetic practice of separating coralline algae into two categories solely on the basis of the presence or absence of genicula does not accurately reflect the evolutionary history of the group.     

Taxonomic contributions to Hapalidiales (Corallinophycidae,Rhodophyta): Boreolithothamnion gen. nov., Lithothamnion redefined and with three new species and Roseolithon with new combinations

Phylogenetic analyses of rbcL gene sequences and of concatenated rbcL, psbA, and nuclear SSU rRNA gene sequences resolved the generitype of Lithothamnion, L. muelleri, in a clade with three other southern Australian species, L. kraftii sp. nov., L. saundersii sp. nov., and L. woelkerlingii sp. nov. Cold water boreal species currently classified in Lithothamnion and whose type specimens have been sequenced are transferred to Boreolithothamnion gen. nov., with B. glaciale comb. nov. as the generitype. The other species are B. giganteum comb. nov., B. phymatodeum comb. nov., and B. sonderi comb. nov., whose type specimens are newly sequenced, and B. lemoineae comb. nov., B. soriferum comb. nov., and B. tophiforme comb. nov., whose type specimens were already sequenced. Based on rbcL sequences from the type specimens of Lithothamnion crispatum, L. indicum, and L. superpositum, each is recognized as a distinct species and transferred to the recently described Roseolithon as R. crispatum comb. nov., R. indicum comb. nov., and R. superpositum com. nov., respectively. To correctly assign species to these three genera based only on morpho-anatomy, specimens must have multiporate conceptacles and some epithallial cells with flared walls. The discussion provides examples demonstrating that only with phylogenetic analyses of DNA sequences can the evolution of morpho-anatomical characters of non-geniculate corallines be understood and applied at the correct taxonomic rank. Finally, phylogenetic analyses of DNA sequences support recognition of the Hapalidiales as a distinct order characterized by having multiporate tetra/bisporangial conceptacles, and not as a suborder of Corallinales whose tetra/bisporangial conceptacles are uniporate.     

Lectotypifícation of Lithophyllum arcticum (Corallinales, Rhodophyta) and a study of its relationships within the Melobesioideae

Three separate collections of the type material of Kjellman's Lithophyllum arcticum are re‐examined and a lectotype is selected. It is confirmed that the thallus is unattached, at least 4.5 cm in diameter, composed of up to eight superimposed more or less discoidal lamellae, provided with concentric striations on the surface. Individual lamellae are usually 100 to 200 μm thick (reaching 1 mm), developing dorsally from the main thallus and expanding centrifugally. The internal organization is dorsiventral with a polystromatic hypothallium, giving rise to an ascending perithallium with small subepithallial initials and rectangular (in TS) epithallial cells. It is found that patches of coaxial‐like growth occur sporadically in the hypothallium and the perithallium, and that ventral lamellae may grow back‐to‐back. Somatic cells exhibit both large and narrow cell fusions. Pore plates of the raised multiporate conceptacles are slightly sunken to flattened and perforated by 16 to 31 pores. Pore canals are conical (narrowing towards the top) and are bordered by filaments composed of both undifferentiated and slender‐elongate cells. Old conceptacles are overgrown by vegetative filaments and empty chambers are embedded in the perithallium. Collectively these features indicate that L. arcticum belongs to the subfamily Melobesioideae. The development of an unattached‐superimposed thallus, patches of coaxial growth, short subepithallial initials and specialized pore cells suggest a position either in Mesophyllum, or in an amended Leptophytum to include even species with coaxial patches and unattached‐superimposed habit (characters presently segregating Leptophytum from Synarthrophytorn). The holotype of Lithophyllum zonatum from East‐Finnmark, previously considered to be related to L. arcticum, is re‐examined and shown to belong to a different species. A previous Arctic record of Mesophyllum lichenoides from Spitsbergen is abolished, and thus the disjunct distribution of I. arcticum in relation to Mesophyllum suggests a position in the Synarthrophyton‐Leptophytum complex which shows a bipolar to temperate distribution.     

REPRODUCTION AND LIFE HISTORY OF THE RED ALGA GALAXAURA OBLONGATA (NEMALIALES,GALAXAURACEAE)1

Culture and morphological studies showed that Galaxaura oblongata (Ellis et Solander) Lamouroux has a triphasic life history with conspicuous gametophytes and small filamentous tetrasporophytes. Development of male and female reproductive structures is very similar and both begin with the enlargement of a terminal cell of a filament branch occupying a normal vegetative position within the apical pit of a thallus branch. In male thalli this modified branch forms a conceptacle in which spermatangia are produced. In female thalli, this modified branch forms a three-celled carpogonial branch consisting of a carpogonium, hypogynous cell and basal cell. Filament branches from the basal cell form a pericarp and the gonimoblast develops directly from the carpogonium. Carposporangia are produced in conceptacles which resemble the male conceptacles. About the time the first carposporangia are produced, the carpogonium, hypogynous cell and basal cell form a large fusion cell. Released carpospores germinate in a unipolar or bipolar manner and form small filamentous thalli. Under short day conditions, cruciate tetrasporangia are produced in small clusters. Tetraspores germinate similarly to carpospores and also form small filamentous thalli. Under low nutrient conditions, small cylindrical thalli develop on the filaments and these appear similar to gametophytes collected in nature.     

Utility of psbA and nSSU for phylogenetic reconstruction in the Corallinales based on New Zealand taxa

A number of molecular studies of the Corallinales, a calcified order of the red algae, have used the conservative nSSU gene to investigate relationships within the order. However interspecific variation at this locus is low for closely related species, limiting resolution of recently diverged groups. In this study, we obtained psbA sequence data from specimens of the order from New Zealand that had been identified according to current taxonomic criteria. We compared phylogenetic analyses based on psbA with those based on nSSU for the same dataset, and also analysed nSSU sequences of the New Zealand material with nSSU sequences of Corallinales taxa from other parts of the world. Our study shows that psbA has considerable potential as a marker for this group, being easily amplified and considerably more variable than nSSU. Combined analyses using both markers provide significant support for relationships at both distal and terminal nodes of the analysis. Our analysis supports the monophyly of all three families currently defined in Corallinales: the Sporolithaceae, Hapalidiaceae and Corallinaceae, and indicates cryptic speciation in Mesophyllum and Spongites.     

REVISION OF THE MASTOPHOROIDEAE (CORALLINALES,RHODOPHYTA) AND POLYPHYLY IN NONGENICULATE SPECIES WIDELY DISTRIBUTED ON PACIFIC CORAL REEFS1

The subfamily Mastophoroideae (Corallinaceae, Rhodophyta) is characterized by species possessing nongeniculate, uniporate tetrasporangial conceptacles without apical plugs, the presence of cell fusions, and the absence of secondary pit connections. However, molecular phylogenetic studies not including the type genus Mastophora indicated that the Mastophoroideae was polyphyletic. Our molecular phylogenetic analysis of the subfamily including the type genus using DNA sequences of SSU rDNA and plastid‐encoded gene of PSII reaction center protein D1 (psbA) revealed that Mastophora formed a robust clade only with Metamastophora. The other mastophoroid genera were divided into six lineages within the family Corallinaceae. Five supported lineages—(i) Pneophyllum; (ii) Hydrolithon gardineri (Foslie) Verheij et Prud’homme, Hydrolithon onkodes (Heydr.) Penrose et Woelk., and Hydrolithon pachydermum (Foslie) J. C. Bailey, J. E. Gabel et Freshwater; (iii) Hydrolithon reinboldii (Weber Bosse et Foslie) Foslie; (iv) Spongites; and (v) Neogoniolithon—were clearly distinguished by the combination of characters including the presence or absence of palisade cells and trichocytes in large, tightly packed horizontal fields and features of tetrasporangial and spermatangial conceptacles. Therefore, we amend the Mastophoroideae to be limited to Mastophora and Metamastophora with a thin thallus with basal filaments comprised of palisade cells, tetrasporangial conceptacles formed by filaments peripheral to fertile areas, and spermatangia derived only from the floor of male conceptacles. This emendation supports Setchell’s (1943) original definition of the Mastophoroideae as having thin thalli. We also propose the establishment of three new subfamilies, Hydrolithoideae subfam. nov. including Hydrolithon, Porolithoideae subfam. nov. including the resurrected genus Porolithon, and Neogoniolithoideae subfam. nov. including Neogoniolithon. Taxonomic revisions of Pneophyllum and Spongites were not made because we did not examine their type species.     

A taxonomic reassessment of Spongites (Corallinaceae,Rhodophyta) based on studies of Kützing's original collections

Results from critical studies of the original collections upon which Spongites Kützing, 1841 is based have led to the designation of S. fruticulosa as lectotype species and to the resurrection and recognition of Spongites as a distinct genus of Corallinaceae (Rhodophyta). Spongites is characterized by the absence of geniculae, uniporate tetrasporangial conceptacles, a multistratose non-palisade and non-coaxial medulla (“hypothallium”) and cortex (“perithallium”), fusions between cells of adjacent filaments, and trichocytes which are solitary or arranged in a vertical series. Since at least 1883, the Kützing epithet “fruticulosa” has been misapplied widely to a taxon with multiporate tetrasporangial conceptacles, whereas the type collection of S. fruticulosa possesses uniporate tetrasporangial conceptacles. Of the six original species, three (S. fruticulosa, S. racemosa, S. stalactitica) are retained in Spongites; S. dentata is referred to Lithophyllum and S. nodosa to Lithothamnion as distinct species; and S. confluens is regarded to be conspecific with Lithophyllum incrustans Philippi. Detailed morphogological-anatomical accounts of specimens in the type collections are presented along with relevant historical data on the genus and on the various species studied. The relationships of Spongites to Neogoniolithon and to other genera of Corallinaceae also are discussed.     

Lithophyllum cuneatum sp. nov. (Corallinaceae, Rhodophyta), a new species of non-geniculate coralline alga semi-endophytic in Hydrolithon onkodes and Neogoniolithon sp. from Fiji, South Pacific

A new species of semi-endophytic coralline alga, Lithophyllum cuneatum (Corallinaceae: Lithophylloideae), is described from Fiji. The species is characterized by a wedge-like thallus that is partially buried in the thallus of the host coralline, Hydrolithon onkodes (Heydrich) Penrose et Woelkerling or occasionally Neogoniolithon sp., and that appears at the surface of the host as a small pustule that is usually paler in color than the host. The thallus consists of erect filaments that are derived from a single cell. The basal cell, when visible, is non-palisade, and areas of bistratose margin are absent. Cells of contiguous erect filaments are joined by secondary pit connections. Epithallial cells are present in 2–3 layers, and individual trichocytes are common. Gametangial plants are dioecious. Male conceptacles have simple spermatangial systems that are confined to the floors of their elliptical chambers. Carposporangial conceptacles contain 5–8 celled gonimoblast filaments that are borne at the margin of a more-or-less discoid fusion cell, and so occupy the periphery of the elliptical conceptacle chambers. Tetrasporangial conceptacles are uniporate, with roofs formed from peripheral filaments, and chambers lack a central columella of sterile filaments. Despite its semi-endophytic nature, haustorial cells are absent, and plastids and pigmentation are present.     

The genera Melyvonnea gen. nov. and Mesophyllum s.s. (Melobesioideae,Corallinales, Rhodophyta) particularly from the central Atlantic Ocean

We propose the new genus Melyvonnea to accommodate species previously included in Mesophyllum having: a) perithallial protuberances that may branch and dominate over the encrusting base, b) monoecious gametophytes with gametangial conceptacles occasionally developed in superimposition, c) spheroid carposporangial chambers (lacking a central pedestal), and d) filaments lining canals of multiporate roofs composed of 3 to 5 cells with distinctively elongate basal cells. The new genus shares with Mesophyllum the development of a predominantly coaxial hypothallium. Melyvonnea presently accommodates three species in the Central Atlantic, viz. the generitype Melyvonnea canariensis (Foslie) comb. nov. from the Canary Islands, Melyvonnea erubescens (Foslie) comb. nov. ( = Mesophyllum incertum; type locality: Bermuda) from the western Atlantic, Melyvonnea aemulans (Foslie & Howe) comb. nov. from Puerto Rico, and one Indo‐Pacific species, Melyvonnea madagascariensis (Foslie) comb. nov. We also emend Mesophyllum Lemoine to encompass Northern Hemisphere species that lack the above apomorphies of Melyvonnea and in addition develop a central pedestal in carposporangial conceptacles (via dissolution of the surrounding cells) with gonimoblasts bending down to fill the empty space. Mesophyllum sensu stricto currently includes six species in the northeast Pacific (M. aleuticum, M. conchatum, M. crassiusculum, M. lamellatum, M. megagastri, M. vancouveriense), two species in the western Atlantic (M. mesomorphum and M. syntrophicum), and three species in the northeast Atlantic and the Mediterranean Sea (M. expansum, M. lichenoides, M. philippii). Gametophytic species of each genus show a mainly disjunct distribution being restricted to the tropics–subtropics (Melyvonnea) and the temperate waters of the Northern Hemisphere (Mesophyllum s.s.). This classification is supported by a consensus of studies of all well‐known species of Mesophyllum sensu Adey (1970), and is based on a phylogenetic analysis of morphological and anatomical characters in addition to molecular evidence.     

Evolutionary history of the Corallinales (Corallinophycidae, Rhodophyta) inferred from nuclear, plastidial and mitochondrial genomes

Systematics of the red algal order Corallinales has a long and convoluted history. In the present study, molecular approaches were used to assess the phylogenetic relationships based on the analyses of two datasets: a large dataset of SSU sequences including mainly sequences from GenBank; and a combined dataset including four molecular markers (two nuclear: SSU, LSU; one plastidial: psbA; and one mitochondrial: COI). Phylogenetic analyses of both datasets re-affirmed the monophyly of the Corallinales as well as the two families (Corallinaceae and Hapalidiaceae) currently recognized within the order. Three of the four subfamilies of the Corallinaceae (Corallinoideae, Lithophylloideae, Metagoniolithoideae) were also resolved as a monophyletic lineage whereas members of the Mastophoroideae were resolved as four distinct lineages. We therefore propose to restrict the Mastophoroideae to the genera Mastophora, Metamastophora, and possibly Lithoporella in the aim of rendering this subfamily monophyletic. In addition, our phylogenies resolved the genus Hydrolithon in two unrelated lineages, one containing the generitype Hydrolithon reinboldii and the second containing Hydrolithon onkodes, which used to be the generitype of the now defunct genus Porolithon. We therefore propose to resurrect the genus Porolithon for the second lineage encompassing those species with primarily monomerous thalli, and trichocyte arrangements in large pustulate horizontal rows. Moreover, our phylogenetic analyses revealed the presence of cryptic diversity in several taxa, shedding light on the need for further studies to better circumscribe species frontiers within the diverse order Corallinales, especially in the genera Mesophyllum and Neogoniolithon.     

Coralline red algae from the Silurian of Gotland indicate that the order Corallinales (Corallinophycidae,Rhodophyta) is much older than previously thought

Aguirrea fluegelii gen. et sp. nov. (Corallinales, Corallinophycidae, Rhodophyta) is described from the mid‐Silurian of Gotland Island, Sweden (Högklint Formation, lower Wenlock). The holotype is of dimerous construction and includes a uniporate conceptacle with a sporangium, thus providing evidence that taxa of the Corallinales/Corallinaceae existed at least 300 million years earlier than previously documented. Aguirrea fluegelii cannot be unequivocally placed in any of seven currently recognized lineages/subfamilies/groups of the Corallinaceae as not all diagnostic characters are preserved, and thus is accorded incertae sedis status within the family Corallinaceae and order Corallinales. Extant evolutionary history studies of Corallinophycidae involving molecular clocks now require updating using new calibration points to take account of the much earlier unequivocal mid‐Silurian record of uniporate conceptacle‐bearing taxa of Corallinales/Corallinaceae as well as the parallel record of Graticula, a genus attributed to the Sporolithales.     

The genus Phymatolithon (Hapalidiaceae,Corallinales, Rhodophyta) in South Africa,including species previously ascribed to Leptophytum

Of the genera within the coralline algal subfamily Melobesioideae, the genera Leptophytum Adey and Phymatolithon Foslie have probably been the most contentious in recent years. In recent publications, the name Leptophytum was used in quotation marks because South African taxa ascribed to this genus had not been formally transferred to another genus or reduced to synonymy. The status and generic disposition of those species (L. acervatum, L. ferox, L. foveatum) have remained unresolved ever since Düwel and Wegeberg (1996) determined from a study of relevant types and other specimens that Leptophytum Adey was a heterotypic synonym of Phymatolithon Foslie. Based on our study of numerous recently collected specimens and of published data on the relevant types, we have concluded that each of the above species previously ascribed to Leptophytum represents a distinct species of Phymatolithon, and that four species (incl. P. repandum) of Phymatolithon are currently known to occur in South Africa.Here we present detailed illustrated accounts of each of the four species, including: new data on male and female/carposporangial conceptacles; ecological and morphological/anatomical comparisons; and a review of the information on the various features used previously to separate Leptophytum and Phymatolithon. Southern African species ascribed to the genus Phymatolithon may be separated from one another in the field by their growth forms, the substrata on which they are generally found, and the colour of living thalli. A key for identifying southern African specimens in the field is included. Our data support the conclusion that the characters upon which Leptophytum is based are unreliable for generic delimitation from Phymatolithon.     

The nature of the ancestral red alga: inferences from a cladistic analysis

A cladistic analysis of the orders of Rhodophyta is presented. Sixteen taxa and 34 characters comprise the data matrix. Included in the analysis are biochemical and ultrastructural features of pigments, cell walls, cell organelles, mitosis and pit connections as well as vegetative and reproductive characters. The traditional recognition of two classes or subclasses, Bangiophycidae and Florideophycidae, is not supported regardless of whether Porphyridiales, Rhodochaetales or Bangiales is designated the outgroup. Florideophycidae, however, appears to be monophyletic with Bangiales as its sister group. Relationships among taxa with one or two plug cap layers, i.e. Acrochaetiales, Palmariales, Corallinales, Nemaliales, Batrachospermales, Gelidiales and Hildenbrandiales are unresolved. Rhodochaetales, Bangiales and possibly Erythropeltidales are monophyletic, but Porphyridiales is polyphyletic. The class Cyanidiophyceae is not recognized and the included genera are considered to be unicellular red algae belonging to Porphyridiales. Taxa that have been proposed as sister groups for red algae, including Cyanobacteria, Cryptophyta, Glaucophyta and Chlorophyta, and Ascomycetes and Basidiomycetes are discussed in relation to the proposed phylogeny of Rhodophyta.